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Cortinarioid Fungi of New Zealand I Karl Soop Cortinarioid Fungi of New Zealand i Cortinarioid Fungi of New Zealand An Iconography and Key Eleventh Revised Edition Introduction Preface This book is intended as one small step towards bridging an obvious gap in the literature on New Zealand fungi, viz. colour illustrations and descriptions of dark-spored agaricoid and gastroid species that grow in native forests. During many study visits to the country I have repeatedly experienced the need for such a publication, a need that stems from the following observations: • The target group of fungi is well represented in New Zealand. During parts of the season it dominates the mycoflora in vast areas of the native forests. • At the same time these species are poorly known. Only a modest number have been named, described, and published, and even fewer have been illustrated in colour. In Europe and other parts of the world many cortinarioid fungi have proven critical as indicators of valuable biotopes and areas worthy of conservation. There is evidence that this also applies to New Zealand, which adds to the incentive to explore the target group and its role within the ecology of the region. It is my hope that the book will inspire resident mycologists to continue studies of this important and fascinating subject. It is evident that much remains to be done and that many new species will be discovered. This book must therefore be seen as provisional in the sense that it will be subject to periodical updates as new data become available. Inevitably, also some names will be provisional, used as working names or as tentative identification, until they are later confirmed or perhaps amended. This edition describes a number of new taxa based on new findings and on ongoing molecular analysis. The repertory of taxa has been substantially expanded to 160 colour photos and descriptions, all of Cortinarius species and varieties. As a consequence of recent phylogenetic research there is a continued emphasis on sections as the taxonomical rank for structuring the genus. Karl Soop Mora, Sweden in December 2017 ii Cortinarioid Fungi of New Zealand Karl Soop Scope Up until Edition 6, this book included various dark-spored genera encountered in New Zealand1. Due to space limitations, and to the emergence of literature covering a wide range of genera [e.g. Johnston & al. 2007, McMulan-Fisher et al. 2014], subsequent editions have been dedicated exclusively to the genus Cortinarius. The Cortinarii studied in this book are circumscribed in several ways: • The species are all native to New Zealand, many are presumed endemic. They grow in forests, predominantly with Nothofagus ss lato, many with Leptospermum or Kunzea. All form ectomycorrhiza with either of the mentioned arboreal genera2. They are all terrestrial, growing in the leaf litter of the forest floor. • The group is naturally limited by the extent of the author's findings, and it is evident that many more Cortinarii exist in the country. Still, it is my contention that most common species within the stated taxonomical and ecological range are covered. In some cases well-documented species that I have not encountered are keyed out without a picture (these are marked with a star). Apart from Cortinarius (Pers.) Gray the studied species belong to the former genera: Rapacea E. Horak Cuphocybe R. Heim Rozites P. Karst. Thaxterogaster Singer which are now synonymised with Cortinarius [Peintner & al. 2001b, 2002b]. These are used to provide tentative group names in the following, pending a revision of the complete infrageneric taxonomy from ongoing phylogenetic research (see below). The taxon Dermocybe Fr. is handled as a subgenus of Cortinarius. The cortinarioid genus Stephanopus M.M. Moser & E. Horak, described from Patagonia, has not been recorded from New Zealand so far. Neither has the white-spored genus Leucocortinarius (Lge) Sing., growing in the Northern Hemisphere (it has, moreover, been shown not to be closely related to Cortinarius). Taxonomical Notes The genus Cortinarius has been extensively studied during the recent decades, where analyses of relevant DNA sequences have played a major part [e.g. Garnica & al. 2003, 2005, 2016, Peintner & al. 2001b, 2002b; see also Frøslev & al. 2005, 2006, Niskanen et al. 2009, 2011a,b, 2013a,b, Liimatainen et al. 2015]. From these genetic studies emerges a picture of early evolution that originated before the break-up of Pangæa, but continued afterwards with frequent genomic exchange, as evidenced by the number of Cortinarius clades that are shared between the Northern and Southern Hemispheres today [Soop & Gasparini 2011]. Their fruit-bodies carry all the principal hallmarks of Cortinarius (see the next chapter), but due to steadily diverging genetic material over eons of time, there are many differences in their detailed anatomy. Indeed, no species has with certainty been shown to grow naturally in the temperate regions of both hemispheres, a fact attributable to a large extent to the corresponding endemicity of host-tree taxa. On the other hand, repeated studies have demonstrated that the genus, taken globally, is without any doubt monophyletic. Independent evolution in the two hemispheres has led to a number of more recent Cortinarius clades. In the South, many species evolved in the Antarctic region during the cretaceous period. During the subsequent break-up of Gondwana, primitive species migrated with their host trees, mainly Nothofagus precursors, and their descendants are now encountered principally in New Zealand, New Caledonia, Tasmania, and Patagonia (South America). The vast majority of these appear to be endemic to either of the regions mentioned with relatively few certain cases of overlap. 1 Cortinarius, Hebeloma, Psathyloma, Pholiota, Descolea, Descomyces, Weraroa, Tympanella, Octaviania, 2 For simplicity, Leptospermum and Kunzea are here and in the following collectively designated “myrtaceous trees”. Also Nothofagus is to be taken ss lato including the new genera Lophozonia and Fuscospora. Karl Soop Cortinarioid Fungi of New Zealand iii From a taxonomical point of view it follows that infrageneric taxa, described from Europe and North America in the last few centuries, are in many cases ill-fitted to accommodate the species that abound in the former Gondwanan region. Traditional subgeneric and section names, all published in the North, must therefore be used with some care. This is all the more important as genetic studies performed on boreal taxa reveal many cases of polyphyly. A Cortinarius taxonomy aimed at the South Pacific has not yet been proposed, let alone one that embraces the world. The traditional taxonomy defines a number of subgenera, mainly these: Cortinarius Myxacium Dermocybe Telamonia Phlegmacium Their circumscription is based on morphology, such as viscidity and hygrophanity, or on the presence of certain metabolites (notably anthraquinonic pigments [cf. Gruber 1975, Gasparini 2004]). The genetic analyses have demonstrated that these characters are often, if not consistently, convergent. They are nevertheless practical for a quick classification, such as the one required by this book. In summary, the key and descriptions that follow use subgenera in the traditional (morphological) sense with the implicit understanding that they do not usually reflect natural affinities. The same is true for some of the other infrageneric groups, to which I have sometimes given names (within quotes) that neither correspond to a natural group, nor carry any nomenclatural status. Where known, detailed taxonomical and phylogenetic information is given in the descriptions, and is then enclosed in square brackets. What to Look for Cortinarioid fungi generally have a cap and a stipe, although the stipe may be rudimentary in the sequestrate taxa (truffles, see Ch. 30). The latter often look like typical Cortinarii when viewed from above in their habitat, but differ by the lack of well-developed gills. Instead, the hymenium consists of a closed structure (gleba) that may be lamellate, looking like poorly developed gills, or be more loculate (chambered). The spores produced by the hymenium are brownish, causing gills or gleba to darken with maturity in all species considered. A microscopic examination reveals that the spores are verrucose, i.e., provided with warts that may be more or less prominent. Most cortinarioid species possess a (universal) veil, i.e., an exterior sheath of hyphæ that protects the young fruitbody during early development. As the mushroom grows, the veil breaks up, forming telltale patterns on the surface of the fruitbody. The abundance, shape, colour, and viscidity of these patterns are often highly significant for species determination. The veil remnants may form scales, squamules, or tufts on the cap, and girdles, bands, ring, or volva on the stipe, or again merely a thin coating. In comparison with European cortinarioid species, members of the corresponding New Zealand mycota exhibit, on average, more glutinous and sometimes more colourful veils. In addition, the fruit-bodies possess an inner, cobwebby veil (cortina) that covers the young gills during development. It later collapses on the upper part of the stipe, where it may stain brown from discharged spores. It is important not to confuse this cortinal zone with velar girdles that may appear further down on the stipe. In some species, notably in the former genus Cuphocybe, the cortina is rudimentary or absent. In the former genus Rozites the cortina is replaced by a membranous inner veil that persists as a collar on the stipe. A macroscopic examination of a cortinarioid collection may include the following characters, used in subsequent descriptions in this book: • Cap: size, hygrophanity, viscidity, colour, surface (cutis) structure, aspect of the margin. Most cortinarioid fungi in New Zealand (less so in Europe) are more or less hygrophanous, i.e., the colour changes as the cap dries, forming streaks or concentric rings in the cutis. A viscid to glutinous cap surface is very common, and the degree of viscidity is an important character. It is therefore desirable to collect the specimens in moist weather, or at least not in a period of draught.
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